BACKGROUND

Oxygen concentrations in both the open ocean and coastal waters have been declining since at least the middle of the 20th century. This oxygen loss, or deoxygenation, is one of the most important changes occurring in an ocean increasingly modified by human activities that have raised temperatures, CO2 levels, and nutrient inputs and have altered the abundances and distributions of marine species. Oxygen is fundamental to biological and biogeochemical processes in the ocean. Its decline can cause major changes in ocean productivity, biodiversity, and biogeochemical cycles. Analyses of direct measurements at sites around the world indicate that oxygen-minimum zones in the open ocean have expanded by several million square kilometers and that hundreds of coastal sites now have oxygen concentrations low enough to limit the distribution and abundance of animal populations and alter the cycling of important nutrients.

ADVANCES

In the open ocean, global warming, which is primarily caused by increased greenhouse gas emissions, is considered the primary cause of ongoing deoxygenation. Numerical models project further oxygen declines during the 21st century, even with ambitious emission reductions. Rising global temperatures decrease oxygen solubility in water, increase the rate of oxygen consumption via respiration, and are predicted to reduce the introduction of oxygen from the atmosphere and surface waters into the ocean interior by increasing stratification and weakening ocean overturning circulation.

In estuaries and other coastal systems strongly influenced by their watershed, oxygen declines have been caused by increased loadings of nutrients (nitrogen and phosphorus) and organic matter, primarily from agriculture; sewage; and the combustion of fossil fuels. In many regions, further increases in nitrogen discharges to coastal waters are projected as human populations and agricultural production rise. Climate change exacerbates oxygen decline in coastal systems through similar mechanisms as those in the open ocean, as well as by increasing nutrient delivery from watersheds that will experience increased precipitation.

Expansion of low-oxygen zones can increase production of N2O, a potent greenhouse gas; reduce eukaryote biodiversity; alter the structure of food webs; and negatively affect food security and livelihoods. Both acidification and increasing temperature are mechanistically linked with the process of deoxygenation and combine with low-oxygen conditions to affect biogeochemical, physiological, and ecological processes. However, an important paradox to consider in predicting large-scale effects of future deoxygenation is that high levels of productivity in nutrient-enriched coastal systems and upwelling areas associated with oxygen-minimum zones also support some of the world’s most prolific fisheries.

OUTLOOK

Major advances have been made toward understanding patterns, drivers, and consequences of ocean deoxygenation, but there is a need to improve predictions at large spatial and temporal scales important to ecosystem services provided by the ocean. Improved numerical models of oceanographic processes that control oxygen depletion and the large-scale influence of altered biogeochemical cycles are needed to better predict the magnitude and spatial patterns of deoxygenation in the open ocean, as well as feedbacks to climate. Developing and verifying the next generation of these models will require increased in situ observations and improved mechanistic understanding on a variety of scales. Models useful for managing nutrient loads can simulate oxygen loss in coastal waters with some skill, but their ability to project future oxygen loss is often hampered by insufficient data and climate model projections on drivers at appropriate temporal and spatial scales. Predicting deoxygenation-induced changes in ecosystem services and human welfare requires scaling effects that are measured on individual organisms to populations, food webs, and fisheries stocks; considering combined effects of deoxygenation and other ocean stressors; and placing an increased research emphasis on developing nations. Reducing the impacts of other stressors may provide some protection to species negatively affected by low-oxygen conditions. Ultimately, though, limiting deoxygenation and its negative effects will necessitate a substantial global decrease in greenhouse gas emissions, as well as reductions in nutrient discharges to coastal waters.